Thursday, November 15, 2018

[Paleontology • 2018] Gordodon kraineri • The Oldest Specialized Tetrapod Herbivore: A New Eupelycosaur from the Permian of New Mexico, USA

Gordodon kraineri 
Lucas, Rinehart & Celeskey, 2018
  DOI: 10.26879/899 

Gordodon kraineri is a new genus and species of edaphosaurid eupelycosaur known from an associated skull, lower jaw and incomplete postcranium found in the early Permian Bursum Formation of Otero County, New Mexico, USA. It has a specialized dental apparatus consisting of large, chisel-like incisors in the front of the jaws separated by a long diastema from relatively short rows of peg-like maxillary and dentary cheek teeth. The dorsal vertebrae of Gordodon have long neural spines that bear numerous, randomly arranged, small, thorn-like tubercles. The tubercles on long neural spines place Gordodon in the Edaphosauridae, and the dental apparatus and distinctive tubercles on the neural spines distinguish it from the other edaphosaurid genera—Edaphosaurus, Glaucosaurus, Lupeosaurus and Ianthasaurus. Gordodon is the oldest known tetrapod herbivore with a dentary diastema, extending the temporal range of that anatomical feature back 95 million years from the Late Triassic. The dental apparatus of Gordodon indicates significantly different modes of ingestion and intraoral transport of vegetable matter than took place in Edaphosaurus and thus represents a marked increase in disparity among edaphosaurids. There were two very early pathways to tetrapod herbivory in edaphosaurid evolution, one toward generalized browsing on high-fiber plant items (Edaphosaurus) and the other (Gordodon) toward more specialized browsing, at least some of it likely on higher nutrient, low fiber plant items. Gordodon shows a surprisingly early specialization of the dental apparatus and indicates how incomplete our knowledge is of edaphosaurid evolution, disparity and diversity.

FIGURE 2. Holotype skull, lower jaw and incomplete postcranium of Gordodon kraineri, NMMNH P-70796, photograph (1) and bone map (2). Scale equals 10 cm.

FIGURE 3. The skull and lower jaw, as preserved, of the holotype of Gordodon kraineri, NMMNH P-70796, in right lateral view, photograph (1) and line drawing (2). 
Anatomical abbreviations are: an = angular; ar = articular; d = dentary; ep = epipterygoid; f = frontal; l = lacrimal; m = maxilla; mtp = mandibular tooth plate; n = nasal; p = parietal; pf = postfrontal; pm = premaxilla; pr = prearticular; prf = prefrontal; pt = pterygoid; q = quadrate; qj = quadratojugal; sa = surangular; sm = septomaxilla; spl = splenial; sq = squamosal; st = supratemporal; t = tabular; v = vomer. Scale equals 1 cm.


SYNAPSIDA Osborn, 1903

Gordodon gen. nov.

Etymology. Gordo, Spanish for “fat,” and Greek odon, “tooth,” in reference to the large (“fat”) teeth at the anterior end of the snout of the holotype. Gordo also is a reference to the city of Alamogordo, near the type locality.

Diagnosis. Gordodon is a medium-sized edaphosaur (presacral length ~1 m) distinguished from the other edaphosaurid genera by: an unique dental apparatus consisting of large chisel-like incisors in the premaxilla and dentary (dentary incisors inferred from empty alveolus) separated by a long diastema from a relatively short row of peg-like maxillary and dentary cheek teeth and tooth plates with small (<1 mm) teeth on the interior surface of the mandible; preorbital skull length subequal to postorbital skull length; a relatively short nasal-maxilla suture; cervical and anterior dorsal vertebrae with relatively gracile centra that are double-keeled ventrally; and cervical and dorsal vertebrae have long neural spines that bear up to 12 small, thorn-like lateral tubercles randomly distributed on each side.

Gordodon kraineri sp. nov.

 Etymology. To honor Karl Krainer for his many contributions to our knowledge of the late Paleozoic geology and paleontology of New Mexico.

Holotype. NMMNH P-70796, incomplete skeleton consisting of the skull, lower jaws, all or parts of 21 vertebrae (five cervical vertebrae, four complete dorsal vertebrae, the neural spines in varying states of completeness of 12 additional dorsal vertebrae), parts of five cervical and five dorsal rib pairs, parts of the right and left clavicles and scapulae and parts of two digits of the manus(?) (Figure 2).

Holotype locality. NMMNH locality 8967, Otero County, New Mexico, USA (Figure 1).

Stratigraphic horizon and age. Lower part of Bursum Formation, early Wolfcampian (early Permian).


 Life restoration of Gordodon kraineri.

Spencer G. Lucas, Larry F. Rinehart and Matthew D. Celeskey. 2018. The Oldest Specialized Tetrapod Herbivore: A New Eupelycosaur from the Permian of New Mexico, USA.   Palaeontologia Electronica. 21.3.39A; 1-42.  DOI: 10.26879/899  

Plain Language Abstract: Gordodon kraineri is a new kind of sail-backed reptile based on an incomplete skeleton found in ~300 million year old rocks in southeastern New Mexico. Gordodon belongs to a family of early herbivorous reptiles, the Edaphosauridae, and has a surprisingly specialized skull and dentition. These skeletal specializations indicate it was a selective browser on plants.

[Entomology • 2018] Paramonovius nightking • A New Genus and Species of An Unusual Australian Winter Bee Fly (Diptera: Bombyliidae) with Discussion on Its Phylogenetic Position

Paramonovius nightking 
Li & Yeates, 2018
photo by 'Jean and Fred'

A new genus of bee fly (Bombyliinae: Bombyliini) is described to place a unique species Paramonovius nightking gen. et sp. nov. The new species is only collected in winter and is only known from a restricted area in Western Australia. Our morphological phylogeny indicates the new genus is sister to Sisyromyia White, 1916 and is similar to the latter in some external characters such as the one‐segmented antennal flagellum with apical stylus, subapex with 3–5 long hairs, open cell r5 and cell br nearly as long as cell bm. However, Paramonovius gen. nov. has dichoptic male eyes, sparse scales on the median abdominal stripe, enlarged claw and pulvillus, as well as a series of autapomorphies in the genitalia. We present a modified key to the Australian genera of Bombyliini to accommodate this new genus. The unusual flight time of this genus may have contributed to its rarity in collections.

Keywords: morphological phylogeny, Paramonovius, taxonomy

Paramonovius nightking Wandoo National Park, Western Australia

photo by 'Jean and Fred'

female Paramonovius nightking 

photo by 'Jean and Fred' 

Genus Paramonovius gen. nov.

Etymology: This generic name is in honour of Dr. Sergei Jacques Paramonov for his significant contribution to Australian dipterology.

Paramonovius nightking sp. nov. 

Etymology: This species is named after the Night King in the American fantasy drama Game of Thrones, because all the specimens were collected in winter and the fly is mostly covered in thick pale pruinescence. The specific name is treated as a noun in apposition.

Xuankun Li and David K. Yeates. 2018. A New Genus and Species of An Unusual Australian Winter Bee Fly (Diptera: Bombyliidae) with Discussion on Its Phylogenetic position. Austral Entomology.  DOI: 10.1111/aen.12361

[Botany • 2019] Didymocarpus middletonii (Gesneriaceae) • A New Species from Limestone Area of central Laos [Nam Kading National Protected Area VI]

Didymocarpus middletonii   Souvann., Soulad. & Tagane

in Souvannakhoummane, Souladeth, Tagane, et al., 2019. 
ດອກລຳໂພງຫີນສີມ່ວງ |

Didymocarpus middletonii Souvann., Soulad. & Tagane, a new species of Gesneriaceae from Nam Kading National Protected Area, is described and illustrated. The new species is morphologically similar to Didymocarpus brevicalyx, D. formosus and D. puhoatensis but distinguished from the three by its fewer-flowered inflorescence, longer pedicel, and urceolate and multicellular eglandular hairy calyx. Based on the latest IUCN criteria, Didymocarpus middletonii is proposed to be Critically Endangered (CR). Our record of Didymocarpus represents a new genus record for the flora of Laos.

Keywords: Biodiversity, Didymocarpus brevicalyx, Didymocarpus formosus, Didymocarpus puhoatensis, Indochina, Laos, new taxa, taxonomy.

Fig. 1. Didymocarpus middletonii Souvann., Soulad. & Tagane.
 A, Habit; B, abaxial surface of lamina; C, inflorescence with flower (lateral view); D, flower (top view); E, dissected corolla, showing stamens and staminodes; F, multicellular glandular and eglandular hairs on pedicel; G, calyx and pistil; H, fertile stamens. Scale bars: A, 4 cm; B, 3 cm; C–E and G, 1 cm; F and H, 4 mm.
Drawn by K. Souvannakhoummane from Tagane et al. L1198 (FOF).

Fig. 2. Didymocarpus middletonii Souvann., Soulad. & Tagane in the type locality.
 A, Habit; B, abaxial surface of lamina; C, flower (front view); D, flower (lateral view). Scale bars: A, 4 cm; B–D, 1 cm.
Photographed by S. Tagane on 29 June 2017.

Didymocarpus middletonii Souvann., Soulad. & Tagane, sp. nov. 

Didymocarpus middletonii is similar to Didymocarpus brevicalyx Nangngam & D.J.Middleton, Didymocarpus formosus Nangngam & D.J.Middleton from Thailand and Didymocarpus puhoatensis X.Hong & F.Wen from Vietnam but differs from these three in having an urceolate calyx (versus campanulate) and a smaller corolla (c.3.4 cm long versus longer than 4.5 cm). 

– Type: Laos, Nam Kading National Protected Area, Bolikhamxai Province, ..., 665 m elevation, 29 vi 2017, Tagane, S., Souladeth, P., Okabe, N., Yang, C.-J. L1198 [fl.] (holo FOF; iso E, HNL, P). 

Distribution. Laos, Bolikhamxai Province (so far known only from the type locality). 

Habitat and ecology. Didymocarpus middletonii grows on shaded rocks, along a stream in semi-evergreen forest. Given that most of the individuals seen during our field survey had flower buds, the species should be in full bloom in July. 

Etymology. The specific epithet honours Dr David Middleton (Singapore Botanic Gardens), from whom we received the most generous advice regarding this new species.

Vernacular name. ດອກລຳໂພງຫີນສີມ່ວງ [Dok Lam Phong Hin Si Mouang (meaning: limestone purple funnel flower)].

K. Souvannakhoummane, P. Souladeth, S. Tagane, C.-J. Yang. and T. Yahara. 2019. Nam Kading National Protected Area VI: Didymocarpus middletonii (Gesneriaceae), A New Species from Limestone. Edinburgh Journal of Botany.  DOI: 10.1017/S0960428618000264

[Botany • 2018] Five New Species of Syzygium (Myrtaceae) from Indochina and Thailand: Syzygium honbaense, S. phamhoangii, S. yersinii, S. phoukhaokhouayense & S. scabrum

ຫວ້າພູ  ||  Syzygium phoukhaokhouayense Soulad., Tagane & Yahara

in Tagane, Dang, Souladeth, Nagamasu, Toyama, et al., 2018. 
 Photographs: S. Tagane

Five new species of Syzygium (Myrtaceae), Syzygium honbaenseSphamhoangii and S. yersinii from Khanh Hoa Province, Vietnam, Sphoukhaokhouayense from Phou Khao Khouay National Protected Area, Vientiane Province, Laos, and S. scabrum from Bung Khla, Phu Wua Wildlife Sanctuary, Buengkan Province, Thailand, are described and illustrated. Photographs, vernacular names and preliminary conservation assessments are provided for them.

Keywords: Flora, Laos, Myrtales, new species, Thailand, taxonomy, Vietnam, Eudicots

Syzygium honbaense Tagane, V.S.Dang & Yahara, sp. nov.
TYPE:— VIETNAM. Khanh Hoa Province, Mt. Hon Ba, ...

Syzygium honbaense is distinct from all the other Syzygium species in the region by having terminal inflorescences with 3 to 5 reddish flowers, ca. 1.1 cm long hypanthium and relatively smaller leaves, to 7.2 × 2.7 cm

Distribution:—Vietnam (so far only known from the type locality, Mt. Hon Ba). 
Habitat and Ecology:—Hill evergreen forest, at ca. 1200 m elevation. 

 Etymology:— The new species is named after the type locality, Hon Ba Nature Reserve in Khanh Hoa Province of Vietnam.
 Vernacular name:— Trâm hòn bà.

Note:— Among the Syzygium species having relatively large (hypanthium > 1 cm in diam.) and reddish to purplish flowers in Indochina, S. honbaense is easily distinguished from the other species by a combination of its smallest leaves ((1.7–)2.6–7.2 cm long vs. longer than (6–)12 cm) and terminal inflorescences. 

Syzygium phamhoangii Tagane, V.S.Dang & Yahara, sp. nov. 
TYPE:—VIETNAM. Khanh Hoa Province, Mt. Hon Ba, ...

Syzygium phamhoangii is similar to Syzygium balsameum (Wight 1841: 16) Walpers (1843: 179) in the shape of leaves and axillary inflorescences but distinguished by its obtuse to slightly cordate leaf base (vs. cuneate to long attenuate), shorter petioles ((1–)2–4 mm long vs. 4–15 mm long), larger hypanthium (3.5–4 mm long vs. 2.5–3.5 mm long) and more ovules per locule in ovary (12–16 ovules vs. 3–8 ovules).

Distribution:—Vietnam (so far only known from the type locality, Mt. Hon Ba). 
Habitat and Ecology:—Hill evergreen forest, at ca. 920 m elevation. 

 Etymology:— The specific epithet is chosen in honor of the excellent Vietnamese botanist Prof. Dr. Phạm Hoàng Hộ, who significantly contributed to the study of the flora of Vietnam. 
Vernacular name:— Trâm phạm hoàng hộ.

Syzygium yersinii Tagane, V.S.Dang & Yahara, sp. nov. 
TYPE:— VIETNAM. Khanh Hoa Province, Mt. Hon Ba, ...

Syzygium yersinii is similar to S. chantaranothaianum Soh & Parnell (2012: 558) in ovate-oblong leaf shape, very short petioles and terminal inflorescences but differs in having larger and thicker leaves (thickly coriaceous in S. yersinii vs. subcoriaceous in S. chantaranothaianum), reticulate tertiary vein (vs. scalariform) and larger flowers (hypanthium 1.8 cm long vs. 0.8–1 cm long).

Distribution:— Vietnam (so far only known from the type locality, Mt. Hon Ba). 
Habitat and Ecology:— Hill evergreen forest, at 890–920 m elevation. 

 Etymology:— This species is named after Dr. Alexandre Emile Jean Yersin (1863–1943), a Swiss-French, for his contributions to the exploration of the Hon Ba mountain. 
 Vernacular name:—Trâm yersin.

Note:— Lee et al. (2014: 398) identified this species as Syzygium formosum (Wallich 1831: 108) Mason (1851: 554) but S. yersinii is easily distinguished from S. formosum by its opposite leaves (vs. usually whorled in S. formosum) and terminal inflorescences (vs. in the axils of fallen leaves). The leaf texture and venation when dry is very similar to S. grande (Wight 1841: 17) Walpers (1843: 180), but differs from S. grande in its small habit (4 m tall vs. usually more than 20 m tall), subsessile leaves (vs. petiolate in S. grande) and more or less slightly cordate leaf base (vs. cuneate).

FIGURE 4. Syzygium phoukhaokhouayense Soulad., Tagane & Yahara. 
A) Flowering branch, B) Portion of lower leaf surface, C) Bark, D) Young shoot, E & F) Flowers. 
Photographs: A–F for Yahara et al. L1827,
 taken by S. Tagane on 26 Dec. 2017.
Syzygium phoukhaokhouayense Soulad., Tagane & Yahara, sp. nov. 
 TYPE:—LAOS. Vientiane Province, Thoulakhom district, Ban Pa Paek, Phou Khao Khouay National Protected Area, ...

Syzygium phoukhaokhouayense is similar to S. syzygioides (Miquel 1855: 431) Merrill & Perry (1938: 109) but differs in having coriaceous leaves (vs. chartaceous to subcoriaceous in S. syzygioides), longer petiole (7–12 mm long vs. 3–5 mm long), larger and reddish-purple hypanthium (4.1–4.5 mm long vs. ca. 3 mm long, greenish), longer styles (ca. 5 mm long vs. ca. 8.3 mm long) and fewer ovules per locule in ovary (4–5 per locule in S. phoukhaokhouayense vs. 10–14 per locule in S. syzygioides). Also, it is apparently similar to S. lineatum (De Candolle 1828: 287) Merrill & Perry (1938: 109) but easily distinguished by having more secondary veins (28–32 pairs in S. phoukhaokhouayense vs. 16–20 pairs in S. lineatum) and single intramarginal veins (vs. 2).

Distribution:—Laos (so far only known from Phou Khao Khouay National Protected Area). 
 Habitat and Ecology:—Open pine forest, at 905 m elevation.  

Etymology:— The species epithet refers to the geographical location of the find, Phou Khao Khouay National Protected Area. 
 Vernacular name:— ຫວ້າພູ (Wa Phou). 

Note:— The matK sequence of S. phoukhaokhouayense is identical with S. syzygioides (783/783 bp for GenBank accession no. AB924771 and AB925281, 780 bp for AB924710 and AB924734, 768/768 bp for AB924947). However, S. phoukhaokhouayense is easily distinguished from S. syzygioides by the diagnostic characters mentioned above, such as the differences in leaf thickness, length of petiole and style, and colour of hypanthium.

FIGURE 5. Syzygium scabrum Tagane, Soulad. & Yahara. 
A) Leafy twig, B) Portion of lower leaf surface, C) Inflorescence, D) Flowers and flower buds, E) Young fruit, F) Fruit and seed. 
Photographs: A­ & B for Yahara et al. L1727, taken by S. Tagane on Dec. 2017; 
C & D for Souladeth 86, taken by P. Sutthisaksopon on 24 May 2011; 
E & F for Phonsena et al. 7280 taken by P. Phonsena on 22 Nov. 2015.

Syzygium scabrum Tagane, Soulad. & Yahara, sp. nov. 
TYPE:—THAILAND. Buengkan Province, Bung Khla, Phu Wua Wildlife Sanctuary, ...

Syzygium scabrum is similar to S. vestitum Merrill & Perry (1938: 110) in having reddish brown hairs on twigs, leaves and hypanthium, but differs in having more or less cordate leaf base (vs. broadly cuneate to rounded in S. vestitum), scabrid on both sides of leaf surfaces (vs. glabrous except on veins on abaxial side, never scabrid on adaxial side), more secondary veins (16–30 pairs vs. 10–16 pair) and longer styles (8–15.5 mm long vs. 6 mm long).

Distribution:— Laos (Vientiane Province: Phou Khao Khouay National Protected Area), Thailand (Nakhon Phanom Province: Phu Langka National Park, Buengkan Province: Phu Wua Wildlife Sanctuary). 
 Habitat and Ecology:—In hill evergreen forest, at altitudes of 690–770 m in Laos, and in dry evergreen forest at an altitude of 150 m in Thailand. 

 Etymology:—The species epithet refer to its nature of roughened (scabrid) surfaces of twigs and leaves caused by dense hairs. 
 Vernacular name:— หว้าขน (Wa Khon) (Thailand); ຫວ້າຂົນ (Wa Khon) (Laos).

Note 1:—This species has been confused with S. vestitum (type: Mt. Bana, Vietnam, J. & M.S. Clemens 3296, K, image!) by Souladeth & Meesawat (2012), Chantaranothai (2014), and Soh & Parnell (2015), but it is clearly distinguished from S. vestitum by the above diagnostic characters. Syzygium vestitum is restricted to northern to central Vietnam and southern China (southeast Yunnan) (Hô 2003, Chen & Craven 2007, from our field observations in SE Asia). The sequence of matK region of S. scabrum (GenBank accession no. LC381853) differs 7 bp of the total 760 bp from the S. vestitum (LC381852: Tagane et al. V2522 (FU!) from Bach Ma National Park, Central Vietnam, ca. 25 km apart from the type locality of S. vestitum), supporting the separation of the two species.


Shuichiro Tagane, Van-Son Dang, Phetlasy Souladeth, Hidetoshi Nagamasu, Hironori Toyama, Akiyo Naiki, Kengo Fuse, Hop Tran , Cheng -Jui Yang, Amornrat Prajaksood and Tetsukazu Yahara. 2018.  Five New Species of Syzygium (Myrtaceae) from Indochina and Thailand.  Phytotaxa. 375(4); 247–260.  DOI: 10.11646/phytotaxa.375.4.1


[Botany • 2018] The True Tectaria chinensis (Tectariaceae): Morphology, Distribution, and Allied Species

Tectaria chinensis (Ching & Chu H. Wang) Christenh.
in Dong, Tan, Pham & Phan, 2018. 

Tectaria chinensis is a poorly known species from China, with its type being the sole authentic voucher to date. Recent field observations and morphological comparisons reveal its type being an incomplete frond, which resulted in the constant misinterpretation of some morphological characters and the allied species to T. chinensis. In fact, T. chinensis is a very special species restricted in southern China and northern Vietnam and featured by the unique long and erect caudex and the rare combination of partly free venation and vein-dorsal sori. Phylogenetically, T. chinensis was revealed by our previous analyses of five plastid regions to be in the Ctenitopsis group (Clade III or T. subg. Ctenitopsis), forming a sister clade to the lineage of T. fuscipes. A detailed description, explanatory illustrations, as well as distribution and habitat information of T. chinensis are provided.

Keywords: China, morphology, taxonomy, Tectaria, Vietnam, Pteridophytes

FIGURE 2. Tectaria chinensis (Ching & Chu H. Wang) Christenh. 
A & B, habit. C, two long, erect caudices. D, scales on lower stipe. E, part of fragment on upper lamina, showing veins forming costal areoles and sori on anastomosing veins or dorsal on free veins.
 (A from Dong 4343 at IBSC, B–E from Dong 4789 at IBSC).

Tectaria chinensis (Ching & Chu H. Wang) Christenhusz (2010: 58). 
Basionym:— Ctenitopsis chinensis Ching & Chu H. Wang (1981: 124). 
Type:— CHINA. Yunnan: Hekou, in 1955, elev. 100 m, Department of Biology, Yunnan University (DBYU) 512 (holotype PE!)

Distribution:—Restricted to southern China and northern Vietnam. In China: southern Yunnan (Hekou, Jinping, Lüchun, Mengla, and Xichou); in Vietnam: Ha Giang (Cao Bo, Vi Xuyen Distr.), Lao Cai (Nam Xe, Van Ban Distr.). This is the first time to confirm T. chinensis in Vietnam. It is also highly expected to find this species in northern Laos.

Note:— The type of Tectaria chinensis, DBYU 512 (PE), is actually an incomplete frond that consists of a broken stipe and a lamina without basal pinnae (Fig. 1). The lowest pinnae seen on the sheet of type specimen was mistakenly regarded as the basal pinnae by Ching & Wang (1981), which leaded them and later authors (Wang 1999, Cheng 2005, Xing et al. 2013) to associate this species with Ctenitopsis subsageniacea sensu Ching (1938: 311) [= T. austrosinensis (Christ 1907: 145) C. Christensen (1934: 177) (Dong 2017)], a species with the basal basiscopic lobes reduced on basal pinnae. Based on the misinterpretation of T. chinensis, we had never associated our new collections featured by long-erect caudices with T. chinensis because in the new collections the basal basiscopic pinnules of basal pinnae are constantly produced (Fig. 2). On the other hand, of all known species in Tectaria from China T. chinensis is the only dubious species which is poorly presented in herbarium; this fact reminded us to connect our new collections featured by remarkable long caudex with T. chinensis. The overall similarity, especially the consistent venation and sori positions between the new collections and the type of T. chinensis makes it sure that our new collections are just the individuals of T. chinensis. Now it is clear that T. chinensis is with the basal basiscopic pinnules being produced on basal pinnae and is not at all similar to T. austrosinensis.

Shi-Yong Dong, Shi-Shi Tan, Van The Pham and Ke Loc Phan. 2018. The True Tectaria chinensis (Tectariaceae): Morphology, Distribution, and Allied Species. Phytotaxa. 376(1); 60–67.  DOI: 10.11646/phytotaxa.376.1.6

Wednesday, November 14, 2018

[PaleoOrnithology • 2018] Mirarce eatoni • The Most Complete Enantiornithine (Aves, Ornithothoraces) from North America and A Phylogenetic Analysis of the Avisauridae

Mirarce eatoni 
Atterholt​, Hutchison & O’Connor, 2018

 Illustration: Brian Engh. 

The most complete known North American enantiornithine was collected in 1992 but never formally described. The so-called “Kaiparowits avisaurid” remains one of the most exceptional Late Cretaceous enantiornithine fossils. We recognize this specimen as a new taxon, Mirarce eatoni (gen. et sp. nov.), and provide a complete anatomical description. We maintain that the specimen is referable to the Avisauridae, a clade previously only known in North America from isolated tarsometatarsi. Information from this specimen helps to clarify evolutionary trends within the Enantiornithes. Its large body size supports previously observed trends toward larger body mass in the Late Cretaceous. However, trends toward increased fusion of compound elements across the clade as a whole are weak compared to the Ornithuromorpha. The new specimen reveals for the first time the presence of remige papillae in the enantiornithines, indicating this feature was evolved in parallel to dromaeosaurids and derived ornithuromorphs. Although morphology of the pygostyle and (to a lesser degree) the coracoid and manus appear to remain fairly static during the 65 million years plus of enantiornithine evolution, by the end of the Mesozoic at least some enantiornithine birds had evolved several features convergent with the Neornithes including a deeply keeled sternum, a narrow furcula with a short hypocleidium, and ulnar quill knobs—all features that indicate refinement of the flight apparatus and increased aerial abilities. We conduct the first cladistic analysis to include all purported avisuarid enantiornithines, recovering an Avisauridae consisting of a dichotomy between North and South American taxa. Based on morphological observations and supported by cladistic analysis, we demonstrate Avisaurus to be paraphyletic and erect a new genus for “A. gloriae,” Gettyia gen. nov.

Figure 2:  Mirarce eatoniA sampling of the best-preserved cervical and thoracic vertebrae, including the axis.
 (A) Axis in lateral view. (B) Axis in dorsal view. (C) Axis in caudal view. (D) Third cervical vertebra in lateral view. (E) Third cervical vertebra in ventral view. (F) Posterior cervical vertebra in lateral view. (G) Posterior cervical vertebra in ventral view. (H) Thoracic vertebra in lateral view. (I) Thoracic vertebra in ventral view. (J) Thoracic vertebra in anterior view.
 Abbreviations: ds, dens; ep, epipophysis; lg, lateral groove; lr, lateral ridge; pap, parapophysis; prz, prezygopophysis; poz, postzygopophysis; ps, posterior shelf; sp, spinous process; vp, ventral process. Scale bar equals one cm. 
Photos: David Strauss.

 Figure 19: A skeletal reconstruction of Mirarce eatoni showing preserved skeletal elements (white).
Illustration: Scott Hartman.

Systematic paleontology
Class AVES Linnaeus, 1758
Subclass ENANTIORNITHES Walker, 1981

Family AVISAURIDAE Brett-Surman and Paul, 1985

Revised diagnosis: Enantiornithine birds with the following unique combination of morphological features: tarsometatarsus with inclined proximal articular surface; strong transverse convexity of the dorsal surface of the mid-shaft of metatarsal III; a distinct plantar projection of the medial rim of the trochlea of metatarsal III (unambiguously supported in our phylogenetic analysis); and a laterally compressed J-shaped metatarsal I (modified from Chiappe (1993)).

Phylogenetic definition: the last common ancestor of Neuquenornis volans and Avisaurus archibaldi plus all its descendants (Chiappe, 1993).

Included genera: Avisaurus (Brett-Surman & Paul, 1985); Soroavisaurus (Chiappe, 1993); Neuquenornis (Chiappe & Calvo, 1994); Intiornis (Novas, Agnolín & Scanferla, 2010); Mirarce (current study); and Gettyia (current study).


Etymology: Named for its spectacular preservation and level of morphological detail (Latin “mirus” for wonderful), and after Arce, winged messenger of the titans in Greek mythology, for the evidence suggesting a refined flight apparatus in this species.

Type species: Mirarce eatoni sp. nov. (by monotypy)

Etymology: The type species is named in honor of Dr. Jeffrey Eaton, for his decades of work contributing to our understanding of the Kaiparowits Formation and the fossils recovered from it.


Holotype: UCMP 139500, a three-dimensional partial skeleton consisting of several cervical and thoracic vertebrae (including the axis), the pygostyle, almost all phalanges from the left pes and several from the right, a complete humerus, femur, and tarsometatarsus, a partial scapula, coracoid, furcula, and tibiotarsus, as well as fragments of the sternum, radius, ulna, carpometacarpus, and manual phalanges (see Table 1 for measurements of select elements).

Type horizon and locality: UCMP locality V93097, Late Cretaceous (late Campanian 76–74.1 Ma; Roberts, Deino & Chan, 2005) Kaiparowits Formation of Grand Staircase-Escalante National Monument in Garfield County, Utah, USA.

Diagnosis. A large, turkey-sized avisaurid (see above diagnosis) enantiornithine (thoracic vertebrae with centrally located parapophyses; pygostyle cranially forked with ventrolateral processes; furcula dorsolaterally excavated; Chiappe & Walker, 2002) with the following autapomorphies: posterior end of sternum weakly flexed caudodorsally, terminating in a small knob; ulnae with remige papillae present; small, deep, circular pit located just craniolateral to the femoral posterior trochanter; small, triangular muscle scar on medial margin of the femoral shaft just distal to the head followed distally by a much larger proximodistally elongate oval; distinct, rugose ridge-like muscle attachment located on the craniomedial margin of the femur a quarter length from the distal end; and tubercle for the m. tibialis cranialis located at the mid-point of the shaft of metatarsal II on the dorsal surface. The new species is further distinguished by the unique combination of the following characters: acrocoracoidal tubercle very weakly developed and medially located; furcula with truncate (untapered) omal tips weakly developed into articular facets and oriented perpendicular to the axis of the rami; ventral projection of the sternal keel proportionately greater than in most other enantiornithines (similar to condition observed in Neuquenornis); acetabulum fully perforate; medial surface of the medial condyle of the tibiotarsus with deep circular excavation; and elongate, slightly raised, flat, oval surface present on the medial edge of the plantar surface of metatarsal II continuous with a weak medial plantar crest.

Figure 20: A reconstruction of living Mirarce eatoni, illustrating the large body size of this taxon.
 Illustration: Brian Engh.

Revised Systematic Paleontology

Etymology: Named in honor of Mike Getty, a great friend, technician, and field paleontologist, who is dearly missed.

GETTYIA GLORIAE (Varricchio & Chiappe, 1995) new comb.

Holotype: MOR 553E/, a three-dimensional tarsometatarsus missing part of metatarsal IV.

Type horizon and locality: Upper Cretaceous (Campanian) Two Medicine Formation, MOR locality TM-068, Glacier County, Montana, USA.

Diagnosis: small avisaurid enantiornithine with the following unique combination of features: dorsal surface of the tarsometatarsus strongly inclined; attachment for the m. tibialis cranialis located beyond the midpoint of the tarsometatarsus; and distal vascular foramen completely closed by metatarsal IV.

Jessie Atterholt​, J. Howard Hutchison and Jingmai K. O’Connor. 2018. The Most Complete Enantiornithine from North America and A Phylogenetic Analysis of the Avisauridae. PeerJ. 6:e5910.  DOI: 10.7717/peerj.5910

[Entomology • 2018] Tropidomantis kawaharai • First Record of A Praying Mantis (Mantodea: Iridopterygidae) in the mid‐Pacific Marquesas Archipelago

Tropidomantis kawaharai  Brannoch

in Brannoch & Svenson, 2018. 

Two praying mantis specimens were collected on the island of Hiva Oa of the Marquesas Archipelago, an isolated island chain located approximately in the centre of the Pacific Ocean. While external morphological features agree with the diagnosis of the genus Tropidomantis, the unique combination of character states reveals that they represent a new species. Furthermore, this is the first record of a praying mantis collected in the Marquesas Archipelago. We describe the morphology of the species, illustrated by high resolution habitus images, figures of diagnostic features and measurement data for Tropidomantis kawaharai sp. nov.

Keywords: Hiva Oa, Marquesas Islands, new record, new species, Tropidomantis kawaharai

Figure 2. Dorsal and ventral habitus images of Tropidomantis kawaharai sp. nov.:
 (a, b) Holotype GSMC006598; (c, d) Paratype GSMC006599.
 Scale bars = 5 mm.  DOI: 10.1111/AEn.12373 

Figure 4  Key morphological features of Tropidomantis kawaharai:
(a) anterior view of head capsule; (b) ventral view of cervical sclerites; (c) dorsal view of pronotum. Dark grey shading = compound eyes and ocelli; light grey shading = cuticular colour markings; crosshatching indicates cuticular depressions; stippling indicates membranous region.

Abbreviations: icar, anterior rim of intercervical sclerite; icpr, posterior rim of intercervical sclerite; ics, intercervical sclerites; job, juxtaocular bulge; lclp, lateral part of lateral cervical sclerites; lcmp, medial part of lateral cervical sclerites; lcs, lateral cervical sclerites; LMP, lateral margin of pronotum; MK, medial keel; vcs, ventral cervical sclerites. 
Scale bars = 5 mm.  DOI: 10.1111/AEn.12373 

Tropidomantis Stål 1877

Tropidomantis kawaharai Brannoch sp. nov.

Etymology: This species is named in honour of Dr Akito Kawahara, who collected the specimens, for his enthusiastic commitment to the success of students in the field of entomology and whose mentorship encouraged the author to pursue a career in systematic entomology.

Sydney K. Brannoch and Gavin J. Svenson. 2018. First Record of A Praying Mantis, Tropidomantis kawaharai Brannoch sp. nov. (Mantodea: Iridopterygidae), in the mid‐Pacific Marquesas Archipelago. Austral Entomology. DOI: 10.1111/aen.12373  

[Botany • 2018] Splitting Echinocactus: Morphological and Molecular Evidence support the Recognition of Homalocephala as A Distinct Genus in the Cacteae

Figure 1. Echinocactus platyacanthus from Querétaro B E. horizonthalonius from Chihuahua C Homalocephala texensis from Chihuahua D H. parryi from Chihuahua E H. polycephala subsp. polycephala from Sonora F Kroenleinia grusonii from Querétaro. Line bar in fruit photographs is 1 mm.

in Vargas-Luna, Hernández-Ledesma, Majure, et al., 2018.

Molecular phylogenetic studies of the six currently accepted species in the genus Echinocactus have partially clarified certain aspects of its phylogeny. Most of the studies lack a complete sampling of Echinocactus and are based only in one source of data. Phylogenetic uncertainties in Echinocactus, such as the recognition of Homalocephala as a different genus from Echinocactus, the exclusion of E. grusonii or the affinities of E. polycephalus, are here resolved. Phylogenetic relationships of Echinocactus were reconstructed with a maximum parsimony, a maximum likelihood and a Bayesian approach including 42 morphological characters, four chloroplast markers (atpB-rbcL, trnH-psbA, trnL-trnF and trnK/matK) and two nuclear genes. The utility of these two nuclear regions related to the betalain cycles (DODA and 5GT) are explored and discussed in relation to their potential as phylogenetic markers. Concatenated analyses with morphological and molecular data sets, plus 13 indels (2847 characters and 26 taxa), show general agreement with previous independent phylogenetic proposals but with strong support in order to propose the recognition of a reduced Echinocactus and the recognition of Homalocephala at the generic level. These results recovered a polyphyletic Echinocactus as currently defined. The here-named HEA clade, recovers the species of Homalocephala, Echinocacuts and Astrophytum as a monophyletic group with strong internal support. The Homalocephala (H. texensis, H. parryi and H. polycephala), was recovered as sister to the Echinocactus clade (E. platyacanthus and E. horizonthalonius), plus the Astrophytum clade. Consequently, we propose here to recognise a monophyletic Echinocactus and a monophyletic Homalocephala as two distinct genera with their own molecular and morphological synapomorphies. The evolution of some morphological characters supporting these clades are discussed, the necessary new taxonomic combinations for Homalocephala are proposed and an identification key for the genera, the species and the subspecies of the HEA clade are presented.

Keywords: Cactaceae, HEA clade, morphological character evolution, North American Deserts

Figure 1. A Echinocactus platyacanthus from Querétaro B E. horizonthalonius from Chihuahua C Homalocephala texensis from Chihuahua D H. parryi from Chihuahua E H. polycephala subsp. polycephala from Sonora F Kroenleinia grusonii from Querétaro. Line bar in fruit photographs is 1 mm.

This is the first phylogenetic study that has evaluated and combined molecular data from chloroplast and nuclear genomes with morphology to test the monophyly of all species and subspecies of Echinocactus currently accepted. Here we reinforce the proposal of excluding Echinocactus grusonii from the genus. Nevertheless, the recognition of Kroenleinia grusonii must be deeply evaluated since phylogenetic relationships of the Ferocactus clade (including K. grusonii, Leuchtenbergia, Stenocactus, Thelocactus and Glandulicactus) are still unresolved. The well-known HEA clade was recovered as monophyletic with strong support. This clade is morphologically and molecularly well defined, suggesting its taxonomic recognition. Our results also support the proposal that Echinocactus, as currently accepted (excluding K. grusonii), should be considered as two independent lineages, the Homalocephala and the Echinocactus clades, each one with its own molecular and morphological diagnostic characters, and each one representing different genera. In this study, all of the analyses recovered E. polycephalus within the Homalocephala clade, supporting its inclusion in this taxon. Here we present the new taxonomic combinations for the species of Homalocephala and an identification key for the genera of the HEA clade and for all of their species and subspecies.

Homalocephala parryi (Engelm.) Vargas & Bárcenas, comb. nov.
Homalocephala polycephala (Engelm. & J.M. Bigelow) Vargas & Bárcenas, comb. nov.
Homalocephala polycephala subsp. xeranthemoides (J.M. Coult.) Vargas & Bárcenas, comb. nov.

 Mario Daniel Vargas-Luna, Patricia Hernández-Ledesma, Lucas Charles Majure, Raúl Puente-Martínez, Héctor Manuel Hernández Macías and Rolando Tenoch Bárcenas Luna. 2018. Splitting Echinocactus: Morphological and Molecular Evidence support the Recognition of Homalocephala as A Distinct Genus in the Cacteae.  PhytoKeys. 111: 31-59.  DOI: 10.3897/phytokeys.111.26856

[Botany • 2018] Bulbophyllum chrysolabium (Orchidaceae, Epidendroideae) • A New Species from Yunnan, China

Bulbophyllum chrysolabium L. Li & D.P. Ye

in Li, Ye & Zeng, 2018. 

Bulbophyllum chrysolabium, a new species belonging to section Racemosae from Yunnan, China is described and illustrated. The species is related to B. orientale and B. morphologorum, but differs by having the following set of characters: obliquely broadly-based triangular petals with a long filiform apex; lip densely glandular papillose and conspicuously ciliolate along margins; lip auricles well developed, narrowly falcate, tapering to a long sharp point at the apex; stelidia subulate and twisted inwards, slightly exceeding operculum. The conservation status of B. chrysolabium is assessed and taxonomic notes are provided.

Keywords: Menglian County, new taxa, section Racemosae, taxonomy

Figure 1. Bulbophyllum chrysolabium.
 A Habit B Flower, lateral view C Flower, frontal view D Dorsal sepal, petals and lateral sepals, adaxial view E Lip, lateral view F Lip, ventral view G Pollinia H Operculum, ventral view I Column, ventral view J Column and lip, lateral view.
 Scale bars: 2 cm (A), 2 mm (B–D), 1 mm (E–F, I–J), 0.2 mm (G–H). 
Drawn by Yun-Xiao Liu.

Figure 2. Bulbophyllum chrysolabium.
A Habitat B Inflorescences C Close-up of inflorescence D Flower, lateral view showing floral bract E Flower, frontal view F Dorsal sepal, petal and lateral sepal, abaxial view G Lip, ventral view H Column and lip, lateral view.
Scale bars, 1 mm (G), 2 mm (D–F, H).

Bulbophyllum chrysolabium L. Li & D.P. Ye, sp. nov.

 Diagnosis: Bulbophyllum chrysolabium is distinguished from all known congeners by having the following unique combination of features: obliquely broadly-based triangular petals with a long filiform apex; lip densely glandular papillose on both sides and conspicuously ciliolate along margins; lip auricles well developed, narrowly falcate, tapering to a long sharp point at the apex; stelidia subulate and twisted inwards, slightly exceeding operculum.

Taxonomic notes: Bulbophyllum chrysolabium appears to be related to B. orientale Seidenf. (Seidenfaden 1979: 138), especially in narrowly falcate lip auricles and twisted stelidia, but differs in distinctly longer floral bracts (almost twice as long as the pedicel and ovary); petals with long filiform apices, a rather smaller lip (ca. 2.8 mm long), significantly glandular-papillose and ciliolate at margins; stelidia slightly exceeding operculum and distinctly longer than column. With respect to filiform petals, B. chrysolabium is also superficially similar to B. morphologorum Kräenzl. (1908: 89), however, the latter have a fat, conical protuberance or callus on the front of the column near its base and scape much longer than rachis. In addition, it has subulate, not twisted stelidia, considerably longer than operculum; lip auricles not falcate, but rather obtuse at the apex. A detailed morphological comparison between B. chrysolabium and its allied species is presented in Table 1.

Distribution and habitat: So far known only from Menglian County in southwest Yunnan Province, China, growing as an epiphyte amongst mosses on the tree trunk near the edge of river in rather exposed circumstances in subtropical evergreen broad-leaved forest.

Etymology: The specific epithet comes from the Ancient Greek word chryso-golden” and the Latin derived labium labellum”, referring to the golden-yellow lip of the type.

 Lin Li, De-Ping Ye and Song-Jun Zeng. 2018. Bulbophyllum chrysolabium (Orchidaceae, Epidendroideae, Malaxideae), A New Species from Yunnan, China. PhytoKeys. 111: 61-68. DOI: 10.3897/phytokeys.111.28136